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Fig. 8 in Three-dimensional micro- and nanostructural characteristics of the scleractinian coral skeleton: A biocalcification proxy

Fig. 8. "Ceratotrochus" magnaghii Cecchini, 1914. A. ZPALH.23/4 in oblique view (A 1); distal septal edge enlarged in A2. A3. "Patches of microcrystals" at growing septal edge (RAF). B. ZPALH.23/5. Polished and etched septum sectioned transversely with dRAF that appear to be com − posed of homogenous "microcrystalline" material, SEM micrograph. C. TL M (C1, C2) and MFM (C3) micrographs of longitudinally sectioned septum ZPALH.23/6 in RAF plane. Organic and mineral phases regularly alternate (grayscale enlargement in C 2); brownish organic dRAF components (C1) stained with acridine orange, fluoresce (C2) with bright−green light. Seemingly homogenous dRAF in transverse section (B), sectioned longitudinally is composed of elongated units spaced ca. 5–7 µm (red arrows in C1). Growth direction of septum indicates black arrow. A. Recent, deep−water specimen, SEAMOUNT 2 cruise, Stat. DW 279, 33°55.60'N, 28°23.70'W, 805 m. B, C. Recent, shallow−water specimen from Mediterranean (Marseille, Riou−Grand Conglu submarine cave), 50 m.

Fig. 1. Stephanocyathuspaliferus Cairns, 1977. ZPALH.23 in Three-dimensional micro- and nanostructural characteristics of the scleractinian coral skeleton: A biocalcification proxy

Fig. 1. Stephanocyathuspaliferus Cairns, 1977. ZPALH.23/1 (originally NMNH 46443 lot). Recent, south of Bonaire, 11°18.8'N, 68°22'W, 384–607 m. Pills sta. P−753. July 26, 1968. A–C. Distal (A), lateral (B), and proximal (C) views of corallum. D, E. SEM of septa and paliform lobes. D. Arrow indicates portion of septum enlarged on E. E. "Patches of microcrystals" (fasciculi of Wise 1972) at the growing septal edge here called the Rapid Acretion Front (RAF).

Fig. 11 in Three-dimensional micro- and nanostructural characteristics of the scleractinian coral skeleton: A biocalcification proxy

Fig. 11. Platygyra daedalea (Ellis and Solander, 1786); ZPALH.23/8 (originally NMNH A.G. Humes collection. Acc. number 274378). Recent, 25 m, north of Ankazo−beravina, near Nosy Be, Madagaskar, August 24, 1967. A. Transverse polished section of septum in TLM; note brownish "calcification centers" (dRAF), regular growth increments of fibers (e.g., in encircled area), and dark brown regions (red arrows) of filaments of endolithic organisms (most likely algae, fungi). B. The same septal fragment as A, stained with acridine orange in MFM view; dRAF exhibit very light, green−yellow fluorescence; higher magnifications (C) show delicate greenish bands matching organic components trapped between successive growth increments of fibers (white arrows). D. Transverse polished section of septum in TLM; dRAF in lower right corner; regular growth increments of fibers (TD) are emphasized by levels of brownish bands. E. SEM micrograph of transverse polished and etched section of septum; fibers adjacent to row of dRAF ("calcification centers") taper regularly at ca. 3 µm matching brownish bands in D.

Fig. 7 in Three-dimensional micro- and nanostructural characteristics of the scleractinian coral skeleton: A biocalcification proxy

Fig. 7. Desmophyllum dianthus (Esper, 1794). ZPALH.23/3. Recent, southern Indian Ocean (NE St. Paul Island), MD50 cruise, Stat. 32/CP 145, 38°40.66'S, 77°35.47'E, 825–1020 m. A, B. Polished and etched septum sectioned transversely (A) and longitudinally in RAF plane (B). dRAF form chain of dCRA ("centers of calcification") (A), which sectioned longitudinally exhibit alternating etching relief (B). dCRA (A) are composed of apparently non−crystalline, "blurry" material. C–E. Transverse polished section of septum in TLM (C, D) and MFM (E) micrographs. In TLM (C, D) organic components differentiate into dark−brown zone of wall and septal dRAF and light−brown, banded zone enclosing dRAF; in MFM (E) they exhibit lighter, green−yellow, and darker, greenish fluorescence, respectively. Contact zone between bundles of fibers (also with apparent fractures) contain organic matter that forms a fluorescent network (E); examples highlighted with red arrows. F, G. Septum longitudinally sectioned in RAF plane in TLM (F) and MFM (G). Minute growth increments (less than 5 µm) appear as "denticulations" on fluorescent bars (G). Growth direction within septum indicated by black arrow.

Fig. 6. Flabellumchunii Marenzeller, 1904. ZPALH.23 in Three-dimensional micro- and nanostructural characteristics of the scleractinian coral skeleton: A biocalcification proxy

Fig. 6. Flabellumchunii Marenzeller, 1904. ZPALH.23/2/4. Locality data as in Fig. 5. A–C. Complementary regions of longitudinally sectioned and polished septum (RAF plane): TLM (A), Back−Scattered Electron mode (BSE) (B), and pseudocolor carbon mapping images acquired on the electron microprobe by wavelength−dispersive techniques (C); black/dark blue equals lower concentration (<150 counts per second) whereas yellow−white equals higher concentrations (>150 c/s). Brownish structures exposed at section surface (A), appear darker in BSE mode (B) as it enhances atomic number contrast; elements with lower atomic numbers appear darker, those with higher atomic numbers appear lighter. BSE darker regions (arrows) match exactly to carbon−enriched regions in WDS x−ray mapping image (C, arrows). D–F. TL M (D, E) and MFM (F) micrographs of longitudinally sectioned septum. Organic and mineral phase of dRAF regularly alternate (grayscale enlargement in E); brownish organic components (D) exposed and stained with acridine orangedye, fluoresce (F) with bright−green light. Growth direction within septum indicated by black arrow.

Fig. 8 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 8. Scleractinian Trochocyathus egeri (White 1879), Upper Campanian to Maastrichtian (Upper Cretaceous), Pierre shale (upper part), Dry Creek, Black Hills (South Dakota, USA); USNM 75221. Polished and etched (formic acid, 1%, 20s) septum with aragonite fibers; two enlargements (A, B). AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (C, E), and deflection (D, F) images of 2×2 µm (C, D) and 500×500 nm (E, F) skeletal surface. Nanograins ca. 40–100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 11. Scleractinian Pachysolenia cylindrica Cuif, 1975 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 11. Scleractinian Pachysolenia cylindrica Cuif, 1975, lower Norian (Upper Triassic), Alakir Çay, Turkey; ZPAL V.31/7. Polished and etched (formic acid, 1%, 20s) pachythecal wall consisted of aragonite fibers; two enlargements (A, B). AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (C, E), and deflection (D, F) images of 2×2 µm (C, D) and 500×500 nm (E, F) skeletal surface. Skeletal regions that show effects of oxidizing solution action (upper part of images) do not exhibit distinct nanogranular pattern. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 15 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 15. Galaxea fascicularis (Linnaeus, 1767), skeleton and calicoblastic layer interface. A. FESEM image (reproduced after Clode and Marshall 2003a: figs. 1, 2) of frozen−hydrated specimen showing: nanogranular structure (B, close−up) of calcareous fibers at their entire length (non−etched state), cross−sections of spindle ectodermal cells with spherical intercellular vesicles, and fibrillar organic matrix (asterisk). C. Close−up of mesh−like, fibrillar organic matrix at skeleton−ectoderm interface with attached small nodular structures (white arrows) that, most likely, correspond to calcium enriched regions indicated by X−ray analysis.

Fig. 1 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 1. Synthetic aragonite crystals. Seven day old cluster of acicular aragonite crystals, overall (A) and close−up (B) views. Two day old cluster of acicular aragonite (C) with growth steps (D). No distinct nanograins are recognizable on crystal surface before and after (E–H) treatment with oxidizing solution. AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (E, G), and deflection (F, H) images of 1×1 µm (E, F) and 500×500 nm (G, H) crystal face. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 7 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 7. Scleractinian Rennensismilia complanata (Goldfuss, 1826), Santonian (Upper Cretaceous), Lower Gosau beds, near Gosau, Austria; USNM 499247. Polished and etched (formic acid, 1%, 20s) septum with bundles of aragonite fibers enveloped by structures with positive etching relief interpreted by Sorauf (1999) as sheaths of proteinaceous matrix (A, C, same skeletal regions in different magnifications); in places skeletal etched fibers show regular discontinuities similar to that of extant zooxanthellates (B). AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (D, F), and deflection (E, G) images of 2×2 µm (D, E) and 500×500 nm (F, G) skeletal surface. Nanograins ca. 50–100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 4 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 4. Calcareous sponge Petrobiona massiliana Vacelet and Lévi, 1958, Recent, Marseille, submarine cave, "Grotte du Figuier", depth 10 m; ZPAL V.31/3. Polished and etched (formic acid, 1%, 20s) basal skeleton (spherulites) with calcite fibers; two enlargements (A, B). AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (C, E), and deflection (D, F) images of 2×2 µm (C, D) and 500×500 nm (E, F) skeletal surface. Nanograins ca. 50–100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 6. Scleractinian Paracyathus cupula Reuss, 1871 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 6. Scleractinian Paracyathus cupula Reuss, 1871, Miocene (Neogene), Korytnica, Holy Cross Mts, Poland; ZPAL V.31/5. Polished and etched (formic acid, 1%, 20s) septum with aragonite fibers; two enlargements (A, B). AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (C, E), and deflection (D, F) images of 2×2 µm (C, D) and 500×500 nm (E, F) skeletal surface. Nanograins ca. 50–100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 16 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 16. Main nanostructural patterns of calcium carbonate crystals and their possible diagenetic pathways. Nanostructural spectrum encompasses: crystals without nanograins (A, based on synthetically produced CaCO3 crystals, Fig. 1); crystals composed entirely of nanograins (C, based on Recent biocrystals formed in hydro−organic gel); and crystals with intermediate nanostructural pattern, having a bumpy texture representing degraded/ fused nanograins (B, based on P. cylindrica skeleton, Fig. 11).

Fig. 3 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 3. Scleractinian Goniastrea retiformis (Lamarck, 1816), Recent, Saipan (Cloud Locality A−12, Northern Mariana Islands, Pacific Ocean); ZPAL V.31/2. Polished and etched (formic acid, 1%, 20s) septum with aragonite fibers in two (A, B) enlargements; note negative relief of etched organic components within fibers. AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (C, E), and deflection (D, F) images of 2×2 µm (C, D) and 500×500 nm (E, F) skeletal surface. Nanograins ca. 50–100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 14 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 14. Thermograms showing direct thermogravimetrical (TG; milligrams), differential thermogravimetrical (DTG; arbitrary units), and differential thermo−analytical (DTA; arbitrary units) curves of synthethic aragonite (A) and three samples with two different nanostructural patterns: Triassic Pachysolenia cylindrica (B) without distinct nanograins; Jurassic Isastraea cf. bernensis (C) and Recent Favia stelligera (C) with well developed nanograins. Assumed amount of intraskeletal hydrated organic components was calculated based on distinct weight loss of 400 mg sample that occurred at ca. 300–450°C.

Fig. 2 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 2. Scleractinian Favia stelligera (Dana, 1846), Recent, Lizard Island (Great Barrier Reef, Pacific Ocean), depth 5–10 m; ZPAL V.31/1 (fragment of colony collected by Ann Budd). Polished and etched (formic acid, 1%, 20s) septum with aragonite fibers in two (A, B) enlargements; note negative relief of etched organic components in dRAF zone (upper A) and between fiber's layers. AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (C, E), and deflection (D, F) images of 2×2 µm (C, D) and 500×500 nm (E, F) skeletal surface. Nanograins ca. 50–100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 5. Stylasterid Adelopora fragilis Cairns, 1991 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 5. Stylasterid Adelopora fragilis Cairns, 1991, Recent, New Caledonia, ORSTOM 5, DW 490, 18°54.9'S/163°24,3'E, depth 230 m; ZPAL V.31/4. Polished and etched (formic acid, 1%, 20s) coenosteum with aragonite fibers; two enlargements (A, B). AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (C, E), and deflection (D, F) images of 2×2 µm (C, D) and 500×500 nm (E, F) skeletal surface. Nanograins ca. 50–100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 9 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 9. Scleractinian Isastraea cf. bernensis Étallon in Thurmann and Étallon, 1864. Oxfordian (Upper Jurassic), Ostromice, western Pomerania, Poland; ZPAL H.IV/303. Polished and etched (formic acid, 1%, 20s) septum with aragonite fibers; two enlargements (A, B). AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (C, E), and deflection (D, F) images of 2×2 µm (C, D) and 500×500 nm (E, F) skeletal surface. Nanograins ca. 60 (commonly 80) –100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 12 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 12. Sparry calcite developed between septa of fossil scleractinian corals. A. ZPAL V.31/8 (lower Maastrichtian, Upper Cretaceous, Lubycza Królewska, Lublin Upland, eastern Poland). B. ZPAL V.31/9 (Carnian, Upper Triassic, Alpe di Specie, Dolomites, Italy). Polished and etched (formic acid, 1%, 20s) ZPAL V.31/8 sparry calcite in two enlargements (A1, A3). Back−Scattered Electron Microscopy image (A2) shows complex history of idividual calcite grain, highlighting zones of different elemental composition: those with elements of lower atomic numbers are darker (core of the grain outlines with arrows), whereas those of higher atomic numbers are lighter (outer part). A4–A7, B1, B2, AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (A4, A6, B1), and deflection (A5, A7, B2) images of 2×2 µm (A4, A5) and 500×500 nm (A6, A7, B1, B2) sample surface. Nanograins on A4–A7 are ca. 60–100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.

Fig. 10 in Nanostructure of biogenic versus abiogenic calcium carbonate crystals

Fig. 10. Tropiastraeid scleractinian, undetermined. Upper Carnian (Upper Triassic), Alpe di Specie, Dolomites, Italy; ZPAL V.31/6. Polished and etched (formic acid, 1%, 20s) septum with aragonite fibers; two enlargements (A, B); fiber's tapering (e.g., arrow in B) is possibly related to original organic matter enrichment zones. AFM (contact mode; buffered pH = 8, ammonium persulfate 1%, 10 min.): height−2D projection (C, E), and deflection (D, F) images of 2×2 µm (C, D) and 500×500 nm (E, F) skeletal surface. Nanograins ca. 60 (commonly 80) –100 nm in diameter. Grayscale bars (left) show z−scale (height) of 2D projection images.